Protein that has a function of maintaining a mutation whereby lateral root formation is blocked and a gene encoding the protein

ABSTRACT

An  Arabidopsis thaliana  double mutant ssl2 slr having a mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, obtained by: treating an  Arabidopsis thaliana  slr dominant mutant (FERM BP-8385), which has no lateral roots, with a mutagen; preparing plants of the next generation of the mutagen-treated slr dominant mutant; and selecting a plant that basically preserves phenotypes of the slr dominant mutant but has lateral roots from the plants of the next generation.

CROSS-REFERENCE TO RELATED APPLICATIONS

[0001] This application is based upon and claims the benefit of priority from the prior Japanese Patent Application No. 2003-147765, filed May 26, 2003, the entire contents of which are incorporated herein by reference.

BACKGROUND OF THE INVENTION

[0002] 1. Field of the Invention

[0003] The present invention relates to a factor which has an influence on the phenotype of plants that have no lateral roots (hereinafter also referred to as “lateral rootless phenotype”). The factor is expected to be applicable to artificial control of lateral root formation in plants. More specifically, the present invention relates to a protein having a function of maintaining the mutation whereby lateral root formation is blocked and a gene encoding the protein.

[0004] 2. Description of the Related Art

[0005] The root of dicotyledon plants consists of a primary root which has grown from a radicle of an embryo after germination and lateral roots which have been branched from the primary root. It is known that auxin as a plant hormone is involved in lateral root formation. The SLR (solitary root) gene of Arabidopsis thaliana is also known as a gene encoding a protein which regulates the effect of auxin on lateral root formation. Further, the solitary-root dominant mutant (hereinafter also referred to as “slr dominant mutant”) is known as Arabidopsis thaliana which has a mutation in the SLR gene and forms no lateral roots (Fukaki et al., Plant J. 2002, 29, 153-168). However, there has been no report of a factor which has an influence on the lateral rootless phenotype of the slr dominant mutant and which is expected to be applicable to artificial control of lateral root formation.

BRIEF SUMMARY OF THE INVENTION

[0006] An object of the present invention is to provide a factor which has an influence on the lateral rootless phenotype of a mutant and which is expected to be applicable to artificial control of lateral root formation in plants. More specifically, an object of the present invention is to provide a protein having a function of maintaining the mutation whereby lateral root formation is blocked and a gene encoding the protein.

[0007] The present invention may provide the following means for solving the above-mentioned objects.

[0008] (1) An Arabidopsis thaliana double mutant ssl2 slr having a mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, obtained by: treating (mutagenizing) an Arabidopsis thaliana slr dominant mutant (FERM BP-8385), which has no lateral roots, with a mutagen; preparing plants of the next generation of the mutagen-treated slr dominant mutant; and selecting a plant that basically preserves phenotypes of the slr dominant mutant but has lateral roots from the plants of the next generation.

[0009] (2) An Arabidopsis thaliana double mutant ssl2 slr, which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant (FERM BP-8385) that has no lateral roots, due to an additional mutation of at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant.

[0010] (3) An Arabidopsis thaliana double mutant ssl2 slr, which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant (FERM BP-8385) that has no lateral roots, due to an additional mutation of the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant, wherein the additional mutation is selected from the group consisting of the following (A) to (D):

[0011] (A) a mutation in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0012] (B) a mutation in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0013] (C) a mutation in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; and

[0014] (D) a mutation in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”.

[0015] (4) A mutant gene having a mutation in at least one base of the SSL2 gene (cDNA) shown in SEQ ID NO: 1, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.

[0016] (5) A mutant gene having a mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.

[0017] (6) A mutant gene of the SSL2 gene (cDNA) selected from the group consisting of the following (a) to (c):

[0018] (a) a mutant gene in which the 566th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”;

[0019] (b) a mutant gene in which the 1005th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”; and

[0020] (c) a mutant gene in which the 901th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”.

[0021] (7) A mutant gene selected from the group consisting of the following (d) to (g):

[0022] (d) a mutant gene in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0023] (e) a mutant gene in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0024] (f) a mutant gene in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; and

[0025] (g) a mutant gene in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”.

[0026] (8) A protein selected from the group consisting of the following (a) and (b):

[0027] (a) a protein comprising the amino acid sequence of SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked; and

[0028] (b) a protein comprising an amino acid sequence of SEQ ID NO: 2, in which one or a few amino acids of the amino acid sequence have been deleted, substituted and/or added and which has a function of maintaining a mutation whereby lateral root formation is blocked.

[0029] (9) A gene encoding a protein selected from the group consisting of the following (a) and (b):

[0030] (a) a protein comprising the amino acid sequence of SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked; and

[0031] (b) a protein comprising an amino acid sequence of SEQ ID NO: 2, in which one or a few amino acids of the amino acid sequence have been deleted, substituted and/or added and which has a function of maintaining a mutation whereby lateral root formation is blocked.

[0032] (10) A gene selected from the group consisting of the following (c) or (d):

[0033] (c) a gene comprising the DNA sequence of SEQ ID NO: 1 and encoding a protein having a function of maintaining a mutation whereby lateral root formation is blocked; and

[0034] (d) a gene comprising a DNA sequence of SEQ ID NO: 1, in which one or a few bases of the DNA sequence have been deleted, substituted and/or added and which encodes a protein having a function of maintaining a mutation whereby lateral root formation is blocked.

[0035] As described above, the present invention provides a protein having a function of maintaining a mutation whereby lateral root formation is blocked and the SSL2 gene encoding the protein. Further, the inventors of the present invention have found that, when the function of the SSL2 gene of the invention is lost in the slr dominant mutant, the slr dominant mutant loses the lateral rootless phenotype and does form lateral roots. Accordingly, it is assumed that the protein encoded by the SSL2 gene is a novel regulating factor of plant root formation, especially lateral root formation. Thus, it is expected that growth of plant roots can be artificially regulated by modifying the function of the aforementioned protein. Specifically, it is expected to facilitate root formation in an herbaceous or woody plant of various types in which lateral roots or adventitious roots are not formed, by modifying the function of an SSL2-homologous gene in the plant.

[0036] Additional objects and advantages of the invention will be set forth in the description which follows, and in part will be obvious from the description, or may be learned by practice of the invention. The objects and advantages of the invention may be realized and obtained by means of the instrumentalities and combinations particularly pointed out hereinafter.

BRIEF DESCRIPTION OF THE SEVERAL VIEWS OF THE DRAWING

[0037]FIG. 1 is a view showing a part of a nucleotide sequence of the SSL2 genomic gene (wild type);

[0038]FIG. 2 is a view showing a part of a nucleotide sequence (continued from FIG. 1) of the SSL2 genomic gene (wild type);

[0039]FIG. 3 is a view showing a part of a nucleotide sequence (continued from FIG. 2) of the SSL2 genomic gene (wild type);

[0040]FIG. 4 is a view showing a part of a nucleotide sequence (continued from FIG. 3) of the SSL2 genomic gene (wild type);

[0041]FIG. 5 is a view showing a part of a nucleotide sequence (continued from FIG. 4) of the SSL2 genomic gene (wild type);

[0042]FIG. 6 is a view showing a part of a nucleotide sequence (continued from FIG. 5) of the SSL2 genomic gene (wild type);

[0043]FIG. 7 is a view showing a part of a nucleotide sequence (continued from FIG. 6) of the SSL2 genomic gene (wild type); and

[0044]FIG. 8 is a view showing a nucleotide sequence of a mutant IAA14 gene.

DETAILED DESCRIPTION OF THE INVENTION

[0045] The present invention will be described in detail hereinafter. It should be noted that descriptions below are provided only for illustrating the present invention and do not restrict the present invention.

[0046] [Arabidopsis thaliana Double Mutant ssl2 slr]

[0047] An Arabidopsis thaliana double mutant ssl2 slr of the present invention (which will be also referred to as “double mutant ssl2 slr” hereinafter) is a double mutant obtained by: treating (mutagenizing) an Arabidopsis thaliana slr dominant mutant, which has no lateral roots, with a mutagen; preparing plants of the next generation of the mutagen-treated slr dominant mutant; and selecting a plant that basically preserves phenotypes of the slr dominant mutant but has lateral roots from the plants of the next generation; wherein the double mutant ssl2 slr has a mutation in at least one base (e.g., one or a few bases) of the SSL2 genomic gene shown in SEQ ID NO: 3.

[0048] In another aspect of the present invention, the double mutant ssl2 slr of the present invention is a double mutant which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant that has no lateral roots, due to an additional mutation in at least one base (e.g., one or a few bases) of the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant.

[0049] Specifically, the double mutant ssl2 slr of the present invention includes a double mutant which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant that has no lateral roots, by having “a mutant gene of the SSL2 genomic gene” described below in the slr dominant mutant.

[0050] In one example, the double mutant ssl2 slr of the present invention is a double mutant which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant that has no lateral roots, by having an additional mutation in the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant, wherein the additional mutation is selected from the group consisting of the following (A) to (D):

[0051] (A) a mutation in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0052] (B) a mutation in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”;

[0053] (C) a mutation in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; and

[0054] (D) a mutation in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”.

[0055] The Arabidopsis thaliana slr dominant-mutant (which will be also referred to as a “slr dominant mutant” hereinafter), which is used for producing the double mutant ssl2 slr of the present invention, shows a lateral rootless phenotype. The slr dominant mutant also exhibits additional phenotypes in which root hairs are hardly formed and the gravitropism of root and hypocotyl is aberrant. The gene which causes the aforementioned phenotypes including the lateral rootless phenotype in the slr dominant mutant, i.e., SLR mutant gene, will be referred to as “mutant IAA14 gene” hereinafter.

[0056] Seeds of the slr dominant mutant have been deposited in the identification name of “solitary-root-1 (Arabidopsis thaliana)” on May 22, 2003, under the International Patent Organism Depositary, National Institute of Advanced Industrial Science and Technology (Tsukuba Central 6, 1-1, Higashi 1-chome, Tsukuba-shi, Ibaraki-ken 305-8566, Japan), pursuant to BUDAPEST TREATY ON THE INTERNATIONAL RECOGNITION OF THE DEPOSIT OF MICROORGANISMS FOR THE PURPOSES OF PATENT PROCEDURE. The accession number “FERM BP-8385” was assigned thereto. The slr dominant mutant can be reproduced by performing self-pollination of the slr dominant mutant, thereby preparing the next generation, and selecting plants that has no lateral roots from the next generation.

[0057] In the production of the double mutant ssl2 slr of the present invention, a slr dominant mutant is at first subjected to a mutagen treatment. As the slr dominant mutant to be subjected to the mutagen treatment, seed, plant body, callus and the like may be used. With regard to the mutagen treatment, known techniques may be employed in the present invention. Specific examples of the mutagen include: a chemical mutagen such as an alkylating agent which alkylates bases of DNA; an electromagnetic wave which causes damage to DNA such as X-rays and ultraviolet rays; and a radioactive substance. Alternatively, the mutagen treatment may be effected according to the known Agrobacterium infection method, in which a DNA region sandwiched between a pair of border sequences (25 base pairs) present at both ends of T-DNA region of Ti plasmid contained in Agrobacterium is inserted into a random site of genome DNA of the slr dominant mutant. Preferably, the mutagen treatment is carried out by immersing seeds of the slr dominant mutant for 12 to 16 hours in a solution containing a chemical mutagen (e.g., ethylmethanesulfonic acid) at a concentration of 0.2 to 0.3% by weight. In the case in which seeds are used as the slr dominant mutant, the slr dominant mutants (seeds) are each grown to plants.

[0058] Next, the slr dominant mutant (plant body) which has been subjected to the mutagen treatment is made to perform self-pollination and the next generation thereof is produced. Among the thus produced next generation, plants which basically preserve the phenotypes of the slr dominant mutant but form lateral roots (i.e., plants having a mutation caused by the mutagen treatment in a homozygous form) are selected. Here, to “basically preserve the phenotypes of the slr dominant mutant” means maintaining all the characteristics of the slr dominant mutant other than the characteristic of not forming lateral roots. Specifically, “the phenotypes of the slr dominant mutant to be preserved” include a characteristic in which root hairs are hardly formed and a characteristic in which the gravitropism of root and hypocotyl is aberrant.

[0059] The plants selected at this stage are new mutants which suppress the lateral rootless phenotype of the slr dominant mutant. There is a possibility that these new mutants include two types of mutants: an “intragenic suppressor mutant” in which an additional mutation has occurred inside a region of the gene (mutant IAA14 gene) which causes the mutation of the slr dominant mutant and an “extragenic suppressor mutant” in which an additional mutation has occurred outside the region of the gene (mutant IAA14 gene) which causes the mutation of the slr dominant mutant. Therefore, it is preferable to confirm that the selected plant does not have an additional mutation inside a region of the mutant IAA14 gene. In other words, it is preferable to confirm that the mutation of the mutant IAA14 gene dose not go back to the normal IAA14 gene in the selected plant. With regard to the details of this confirmation, the descriptions of examples described below may be referred to. The information on the nucleotide sequence of the mutant IAA14 gene is available from SEQ ID NO: 4 and FIG. 8. In FIG. 8, the exon portions are indicated by capital letters and the intron portions are indicated by small letters.

[0060] The plant selected as described above is the “double mutant ssl2 slr” of the present invention. The double mutant ssl2 slr of the present invention is a double mutant having two mutations: “a slr dominant mutation (originally contained in the slr dominant mutant)” and “a ssl2 recessive mutation (newly caused by the mutagen treatment in the present invention)”. In the present invention, four types of lines (ssl2-1, ssl2-2, ssl2-3 and ssl2-4) were selected as the double mutant ssl2 slr. It has been found that all of the four types of lines have an additional mutation inside the region of the same gene (which will be referred to as “SSL2 genomic gene” hereinafter).

[0061] Accordingly, the production of the double mutant ssl2 slr of the present invention is reproducible as described below. That is, plants which form lateral roots are selected from the next generation of the slr dominant mutants which have been subjected to a mutagen treatment; and it is confirmed that the selected plants do not have an additional mutation inside the region of the mutant IAA14 gene and that the selected plants have an additional mutation in the nucleotide sequence of the SSL2 genomic gene. With regard to the technique by which the mutation in the SSL2 genomic gene is confirmed, the descriptions of examples described below may be referred to.

[0062] Any of the double mutants ssl2 slr of the present invention exhibit at least some recovery of the phenotypes of the slr dominant mutant. That is, in any of the double mutants ssl2 slr, the lateral rootless phenotype are recovered to form lateral roots, but no recovery is observed in the other phenotypes of the slr dominant mutant (i.e., aberration of root hair formation and aberration of gravitropism). From this fact, it is assumed that the gene (SSL2 genomic gene) which has been mutated in the double mutant ssl2 slr of the present invention genetically interacts with the mutant gene (mutant IAA14 gene) of the slr dominant mutant.

[0063] The double mutant ssl2 slr can be reproduced by performing self-pollination of the double mutant ssl2 slr, thereby preparing seeds of the next generation. However, a large number of the seeds of the next generation are not stably obtained, because of the undesirable characteristics of the reproductive organs of the double mutant.

[0064] [SSL2 Gene and Protein Encoded by SSL2 Gene]

[0065] The gene (SSL2 genomic gene) which has been mutated in the double mutant ssl2 slr of the present invention has been identified as At2g25170 gene, according to the mutation map-based cloning for Arabidopsis thaliana. The genetic information on At2g25170 gene is available from the following web page: http://mips.gsf.de/cgi-bin/proj/thal/search_gene? code=At2g25170. It has been confirmed by the present invention that the information on the nucleotide sequence, which was available from the aforementioned web page at the time of filing the present application, is correct but the information from the same source on exon and intron includes errors. Specifically, the inventors of the present invention isolated cDNA of the SSL2 gene which has been mutated in the double mutant ssl2 slr, confirmed the nucleotide sequence thereof, and revealed the correct exon and intron structures of the SSL2 genomic gene (refer to FIGS. 1 to 7).

[0066] The nucleotide sequence of the SSL2 gene (cDNA) is shown in SEQ ID NO: 1 and the amino acid sequence of a protein encoded by the SSL2 gene (cDNA) is shown in SEQ. ID NO: 2. The nucleotide sequence of the SSL2 genomic gene is shown SEQ ID NO: 3. Any nucleotide sequence indicates those not having mutation. The nucleotide sequence of the SSL2 genomic gene (SEQ ID NO: 3) is also shown in FIGS. 1 to 7. In FIGS. 1 to 7, the exon portions are indicated by capital letters and the intron portions are indicated by small letters.

[0067] As a result of a mutation of the SSL2 genomic gene, the slr dominant mutant becomes to form lateral roots. In other words, the normal SSL2 gene is essential for maintaining mutation whereby lateral root formation is blocked in the slr dominant mutant. Thus, it has been revealed for the first time, by the present invention, that the SSL2 gene encodes a protein having a function of maintaining a mutation whereby lateral root formation is blocked.

[0068] Accordingly, the present invention provides a gene comprising the DNA sequence shown in SEQ ID NO: 1 and encoding a protein having a function of maintaining a mutation whereby lateral root formation is blocked. In this gene, one base or a few bases in the DNA sequence shown in SEQ ID NO: 1 may be deleted, substituted and/or added, as long as the gene encodes a protein having a function of maintaining a mutation whereby lateral root formation is blocked.

[0069] Further, the present invention provides a gene encoding the following protein: a protein comprising the amino acid sequence shown in SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked. Regarding this gene, one amino acid or a few amino acids in the amino acid sequence of the aforementioned protein may be deleted, substituted and/or added, as long as the gene encodes a protein having a function of maintaining a mutation whereby lateral root formation is blocked.

[0070] Yet further, the present invention provides a protein comprising the amino acid sequence shown in SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked. Regarding this protein, one amino acid or a few amino acids in the amino acid sequence shown in SEQ ID NO: 2 may be deleted, substituted and/or added, as long as the protein has a function of maintaining a mutation whereby lateral root formation is blocked.

[0071] In addition, in the present invention, it has been revealed that the SSL2 gene encodes a protein homologous with an animal protein “Chromodomain-helicase-DNA-binding 3 (CHD3)” which is involved in the conversion of chromatin structure of a chromosome. No study has been reported of the relationship between lateral root formation and conversion of chromatin structure. It has been, for the first time in the present invention, suggested that the conversion of chromatin structure is involved in lateral root formation.

[0072] [Mutant Gene of SSL2 Gene]

[0073] The “mutant gene of the SSL2 gene (cDNA)” of the present invention is a mutant gene having a mutation in at least one base of the SSL2 gene (cDNA) shown in SEQ ID NO: 1, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.

[0074] In the “mutant gene of the SSL2 gene (cDNA)”, “mutation” represents, for example, substitution, deletion, or addition of at least one base, which mutation has an influence on the phenotype of a mutant that has no lateral roots. In other words, the expression of the mutant gene having the above-described mutation enables the phenotype of a mutant that has no lateral roots to be recovered.

[0075] Specifically, the “mutant gene of the SSL2 gene (cDNA)” of the present invention includes the following mutant genes. However, it should be noted that the “mutant gene of the SSL2 gene (cDNA)” of the present invention is not limited to these specific examples:

[0076] 1) a mutant gene in which at least one base (e.g., one base or a few bases) of the SSL2 gene (cDNA) has been substituted with base(s) of other type(s), whereby a codon designating an amino acid has been replaced with a termination codon; and

[0077] 2) a mutant gene in which at least one base (e.g., one base or a few bases) of the SSL2 gene (cDNA) has been substituted with base(s) of other type(s), whereby a codon designating an amino acid of one type has been replaced with a codon designating an amino acid of another type.

[0078] More specifically, the “mutant gene of the SSL2 gene (cDNA)” possessed by the double mutant ssl2 slr selected in the present invention includes the following mutant genes. The mutant genes (a) to (c) are derived from the lines ssl2-1, ssl2-3, ssl2-4 of the double mutant ssl2 slr, respectively:

[0079] a) a mutant gene in which the 566th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”. As a result of this mutation, tryptophan (TGG) as the 189th amino acid in SEQ ID NO: 2 has been replaced with the termination codon (TAG);

[0080] b) a mutant gene in which the 1005th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”. As a result of this mutation, tryptophan (TGG) as the 335th amino acid in SEQ ID NO: 2 has been replaced with the termination codon (TAG);

[0081] c) a mutant gene in which the 901th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”. As a result of this mutation, glycine (GGA) as the 301th amino acid in SEQ ID NO: 2 has been replaced with asparagine (AGA).

[0082] Further, the “mutant gene of the SSL2 genomic gene” of the present invention is a mutant gene having mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.

[0083] Specifically, the “mutant gene of the SSL2 genomic gene” of the present invention includes the following mutant genes. It should be noted that the “mutant gene of the SSL2 genomic gene” of the present invention is not limited to these specific examples:

[0084] 3) a mutant gene in which at least one base (e.g., one base or a few bases) of the exon portion of the SSL2 genomic gene has been substituted with base(s) of other type(s), whereby a codon designating an amino acid has been replaced with a termination codon;

[0085] 4) a mutant gene in which at least one base (e.g., one base or a few bases) of the exon portion of the SSL2 genomic gene has been substituted with base(s) of other type(s), whereby a codon designating an amino acid of one type has been replaced with a codon designating an amino acid of another type; and

[0086] 5) a mutant gene in which at least one base (e.g., one base or a few bases) of a splice site of the SSL2 genomic gene has been substituted with base(s) of other type(s), whereby an intron of the SSL2 genomic gene has not been excised in the normal manner.

[0087] In the mutant gene, a “splice site” represents a boundary site between exon and intron, i.e., a site at which excision of an intron and recombination of the two exons adjacent to both ends of the intron are carried out during a splicing reaction, any substitution of a base at which splice site disturbs the splicing reaction. Specifically, a splice site includes the donor splice site located at the 5′ end of an intron and the acceptor splice site located at the 3′ end of an intron. Specific examples of the splice site include the conserved sequence “gt” located at the 5′ end of an intron and the conserved sequence “ag” located at the 3′ end of an intron.

[0088] More specifically, the “mutant gene of the SSL2 genomic gene” possessed by the double mutant ssl2 slr selected in the present invention includes the following mutant genes. The mutant genes (d) to (g) are derived from the lines ssl2-1, ssl2-2, ssl2-3, ssl2-4 of the double mutant ssl2 slr, respectively:

[0089] d) a mutant gene in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”. As a result of this mutation at the exon portion of the SSL2 genomic gene, tryptophan (TGG) as the 189th amino acid in SEQ ID NO: 2 has been replaced with the termination codon (TAG);

[0090] e) a mutant gene in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”. As a result of this mutation at the splice site of the SSL2 genomic gene, the intron of the SSL2 genomic gene has not been excised in the normal manner and thus a normal mRNA is not produced;

[0091] f) a mutant gene in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”. As a result of this mutation at the exon portion of the SSL2 genomic gene, tryptophan (TGG) as the 335th amino acid in SEQ ID NO: 2 has been replaced with the termination codon (TAG);

[0092] g) a mutant gene in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”. As a result of this mutation at the exon portion of the SSL2 genomic gene, glycine (GGA) as the 301th amino acid in SEQ ID NO: 2 has been replaced with asparagine (AGA).

EXAMPLES

[0093] Hereinafter, the present invention will be described in detail by examples. It should be noted that the present invention is not limited to the descriptions of these examples.

Example 1 Production of a Double Mutant ssl2 slr Having ssl2 Recessive Mutation and slr Dominant Mutation

[0094] About 5000 seeds of the slr dominant mutant (FERM BP-8385), which forms no lateral roots, were subjected to a mutagen treatment in which the seeds were immersed in 0.2% ethylmethanesulfonic acid (EMS) solution for 16 hours. The mutagen-treated seeds (M1 seeds) were each grown to plants, and self-pollination of the grown plants was performed, thereby preparing the next generation. Among the next generation (30,000 plants), plants which formed lateral roots although they basically maintained the phenotypes of the slr dominant mutant were selected. After confirming that the aforementioned phenotypes of the selected plants was reliably inherited to the next generation of the selected plants, the selected plants were identified as double mutant and named “double mutant ssl2 slr”. In the present example, double mutants ssl2 slr of four lines (i.e., ssl2-1, ssl2-2, ssl2-3 and ssl2-4) were obtained.

[0095] Further, it was confirmed on the basis of the nucleotide sequence that the genome DNA of the double mutant ssl2 slr did not have any additional mutation in a region of the gene (mutant IAA14 gene) causing the slr dominant mutation. That is, it was confirmed that the double mutant ssl2 slr did not correspond to an “intragenic suppressor mutant” in which an additional mutation has occurred inside a region of the mutant IAA14 gene. This confirmation was carried out by amplifying the genomic region including the mutant IAA14 gene, by using the PCR primers shown below. The PCR primer sequences for amplifying the genomic region (1476 base pairs) including the mutant IAA14 gene will be described hereinbelow. IAA14-F1: 5-CATATTCTGATTTAAGACATA-3 (SEQ ID NO: 5) IAA14-R1: 5-AATCAATGCATATTGTCCTCT-3 (SEQ ID NO: 6)

[0096] The following primers were used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. IAA14-F2: 5-TTATGGCTAATCAGAAGAGCG-3 (SEQ ID NO: 7) IAA14-F3: 5-TATTCTCTAAACAAAAAAAAC-3 (SEQ ID NO: 8)

[0097] Further, it was confirmed from the nucleotide sequence that the genome DNA of the double mutant ssl2 slr had a mutation in a region of the SSL2 gene. This confirmation was carried out by amplifying the region of the SSL2 gene, by using primers shown below. Specifically, the nucleotide sequence of the SSL2 gene region was determined by: allotting the SSL2 gene region (9353 base pairs) into 7 sub-regions (A to G); effecting amplification by PCR in each of the sub-regions; determining the entire nucleotide sequence of each PCR product; and comparing the entire nucleotide sequence of each PCR product with the genome DNA sequence of the SSL2 gene of the wild type. If any mutation is found, the plant having the mutation is an ssl2 mutant.

[0098] The PCR primer sequences for amplifying each sub-region (A to G) of the SSL2 gene region were as follows.

[0099] 1) PCR primer sequences for amplifying the sub-region (A) SSL2-F1: 5-aattcgacttctgggtactca-3 (SEQ ID NO: 9) SSL2-R1: 5-AAATTAAGTCCCTCAAGCTGG-3 (SEQ ID NO: 10)

[0100] The following primers were used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F2: 5-actctgaatttgtagAAAGAA-3 (SEQ ID NO: 11) SSL2-F3: 5-GAAGATGATTTTGTTGCCATA-3 (SEQ ID NO: 12)

[0101] 2) PCR primer sequences for amplifying the sub-region (B) SSL2-F4: 5-AAGATGGGGAGCTGGAATATC-3 (SEQ ID NO: 13) SSL2-R2: 5-GGCTCAACACCCTCTAGCATA-3 (SEQ ID NO: 14)

[0102] The following primers were used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F5: 5-CATCCATACCAGCTTGAGGGA-3 (SEQ ID NO: 15) SSL2-F6: 5-CAAGTTTGATGTCCTCCTCAC-3 (SEQ ID NO: 16)

[0103] 3) PCR primer sequences for amplifying the sub-region (C) SSL2-F7: 5-ACATGCCCCCCAAAAAGGAGC-3 (SEQ ID NO: 17) SSL2-R3: 5-CCATCAATTCGCTCGTACTGC-3 (SEQ ID NO: 18)

[0104] The following primer was used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F8: 5-atgtgctgaaactgtgtgtac-3 (SEQ ID NO: 19)

[0105] 4) PCR primer sequences for amplifying the sub-region (D) SSL2-F9: 5-ccattgcttttgctgacgcat-3 (SEQ ID NO: 20) SSL2-R4: 5-ttcgatagccaaccacagtct-3 (SEQ ID NO: 21)

[0106] The following primer was used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F10: 5-ggcatgcaatatgggtgg (SEQ ID NO: 22) cgt-3

[0107] 5) PCR primer sequences for amplifying the sub-region (E) SSL2-F11: 5-TCAGGTATGGATCAAAGG (SEQ ID NO: 23) AGC-3 SSL2-R5: 5-CTCCCCTCACCTTCCATC (SEQ ID NO: 24) AAC-3

[0108] The following primers were used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F12: 5-gtgcacaatcttgtcaaa (SEQ ID NO: 25) tca-3 SSL2-F13: 5-GAGGCACAGAGAGTCGCT (SEQ ID NO: 26) GCT-3

[0109] 6) PCR primer sequences for amplifying the sub-region (F) SSL2-F14: 5-tatacattggtttggtct (SEQ ID NO: 27) gcc-3 SSL2-R6: 5-GTAGGGATAGATGATGAG (SEQ ID NO: 28) CCA-3

[0110] The following primers were used, in addition to the above-described two primers, in order to determine the entire nucleotide sequence of the PCR product. SSL2-F15: 5-ccccgatgcatctaaatt (SEQ ID NO: 29) atc-3 SSL2-F16: 5-ACTAGTTCAGGAGAAGgt (SEQ ID NO: 30) gag-3

[0111] 7) PCR primer sequences for amplifying the sub-region (G) SSL2-F17: 5-ACATGCAGAGACGACTTG (SEQ ID NO: 31) TTG-3 SSL2-R7: 5-cggacttcatcgaaccta (SEQ ID NO: 32) ttc-3

[0112] The above-described two primers were used in order to determine the entire nucleotide sequence of the PCR product.

Example 2 Isolation of SSL2 Gene

[0113] The double mutant ssl2 slr (ecotype: Columbia) having both ssl2 recessive mutation and slr dominant mutation, prepared in Example 1, was crossed with the wild type (ecotype: Landsberg erecta), whereby F1 generation was obtained. Then, F2 generation as the next generation of F1 generation was prepared by performing self-pollination of the F1 generation. By using the genomic DNA of the F2 generation, detailed mapping of the SSL2 gene locus was carried out on the basis of the genomic information of Arabidopsis thaliana. From the result of the mapping, it was found out that the ssl2 recessive mutation is located in a genomic region including 17 genes from gene At2g25140 to gene At2g25300 on the second chromosome.

[0114] Next, in the genomic DNA of the selected four lines (ssl2-1, ssl2-2, ssl2-3 and ssl2-4) of the double mutant ssl2 slr, the nucleotide sequences of the above-described 17 candidate genes were examined. As a result, in all of the four lines of the double mutant ssl2 slr, mutation which presumably causes, the protein encoded by At2g25170 gene, to lose the function thereof was found. On the basis of this discovery, the At2g25170 gene was identified as the SSL2 gene.

Example 3 Experiment in which it was Confirmed that the SSL2 Gene is Involved in the Blocking of Lateral Root Formation

[0115] The slr dominant mutant (FERM BP-8385) is a gain-of-function mutant of the IAA14 gene encoding an auxin-inducible protein, and lateral root formation thereof is completely blocked under normal growth conditions on an agar medium. However, the double mutant ssl2 slr which has both ssl2 recessive mutation and slr dominant mutation, newly prepared in the present invention, formed lateral roots under the same normal conditions, although the formation of lateral roots was not so vigorous as in the wild type. From this result, it was proved that the normal SSL2 gene is essential for maintaining the lateral rootless phenotype (i.e., the blocking of lateral root formation) observed in the slr dominant mutant.

[0116] Additional advantages and modifications will readily occur to those skilled in the art. Therefore, the invention in its broader aspects is not limited to the specific details and representative embodiments shown and described herein. Accordingly, various modifications may be made without departing from the spirit or scope of the general inventive concept as defined by the appended claims and their equivalents.

1 32 1 4155 DNA Arabidopsis thaliana CDS (1)..(4152) 1 atg agt agt ttg gtg gag agg ctt cgc ata cga tct gat agg aaa cca 48 Met Ser Ser Leu Val Glu Arg Leu Arg Ile Arg Ser Asp Arg Lys Pro 1 5 10 15 gtt tat aac cta gat gat tct gat gat gac gac ttc gtt cct aaa aaa 96 Val Tyr Asn Leu Asp Asp Ser Asp Asp Asp Asp Phe Val Pro Lys Lys 20 25 30 gat cga acc ttt gag caa gtc gag gct att gtc aga act gat gcg aaa 144 Asp Arg Thr Phe Glu Gln Val Glu Ala Ile Val Arg Thr Asp Ala Lys 35 40 45 gaa aat gca tgt cag gct tgt ggg gaa agt act aat ctt gta agc tgc 192 Glu Asn Ala Cys Gln Ala Cys Gly Glu Ser Thr Asn Leu Val Ser Cys 50 55 60 aat aca tgc act tat gcg ttc cat gct aaa tgc tta gtt cca cct ctt 240 Asn Thr Cys Thr Tyr Ala Phe His Ala Lys Cys Leu Val Pro Pro Leu 65 70 75 80 aaa gat gct tcc gtg gaa aat tgg aga tgc cct gaa tgt gtt agt cct 288 Lys Asp Ala Ser Val Glu Asn Trp Arg Cys Pro Glu Cys Val Ser Pro 85 90 95 ctt aac gag ata gat aag ata ttg gat tgt gaa atg cgt cct aca aaa 336 Leu Asn Glu Ile Asp Lys Ile Leu Asp Cys Glu Met Arg Pro Thr Lys 100 105 110 tct agt gaa caa ggt tcc tcc gat gcg gaa ccg aag cca att ttt gtg 384 Ser Ser Glu Gln Gly Ser Ser Asp Ala Glu Pro Lys Pro Ile Phe Val 115 120 125 aaa cag tat ctc gtg aag tgg aag gga tta tca tac ctt cac tgc tct 432 Lys Gln Tyr Leu Val Lys Trp Lys Gly Leu Ser Tyr Leu His Cys Ser 130 135 140 tgg gtg cct gag aag gag ttc cag aag gct tat aag tca aat cat cgt 480 Trp Val Pro Glu Lys Glu Phe Gln Lys Ala Tyr Lys Ser Asn His Arg 145 150 155 160 tta aaa acc aga gtg aac aat ttt cac cgt caa atg gag tca ttc aat 528 Leu Lys Thr Arg Val Asn Asn Phe His Arg Gln Met Glu Ser Phe Asn 165 170 175 aac agc gaa gat gat ttt gtt gcc ata cgt cct gag tgg acc act gtt 576 Asn Ser Glu Asp Asp Phe Val Ala Ile Arg Pro Glu Trp Thr Thr Val 180 185 190 gat cgg att ctt gcc tgc aga gag gaa gat ggg gag ctg gaa tat ctt 624 Asp Arg Ile Leu Ala Cys Arg Glu Glu Asp Gly Glu Leu Glu Tyr Leu 195 200 205 gtc aaa tat aaa gag cta tcc tat gat gaa tgt tat tgg gag tca gaa 672 Val Lys Tyr Lys Glu Leu Ser Tyr Asp Glu Cys Tyr Trp Glu Ser Glu 210 215 220 tca gac atc tca acc ttc cag aat gaa att caa agg ttc aag gat gta 720 Ser Asp Ile Ser Thr Phe Gln Asn Glu Ile Gln Arg Phe Lys Asp Val 225 230 235 240 aat tct aga act cgc aga agt aaa gat gtt gac cat aaa aga aat ccc 768 Asn Ser Arg Thr Arg Arg Ser Lys Asp Val Asp His Lys Arg Asn Pro 245 250 255 aga gac ttt caa cag ttt gat cat act cct gaa ttc ctc aaa ggc ttg 816 Arg Asp Phe Gln Gln Phe Asp His Thr Pro Glu Phe Leu Lys Gly Leu 260 265 270 tta cat cca tac cag ctt gag gga ctt aat ttt ttg cgg ttc tcg tgg 864 Leu His Pro Tyr Gln Leu Glu Gly Leu Asn Phe Leu Arg Phe Ser Trp 275 280 285 tca aaa cag acg cat gta atc ctt gct gat gaa atg gga cta ggc aag 912 Ser Lys Gln Thr His Val Ile Leu Ala Asp Glu Met Gly Leu Gly Lys 290 295 300 aca att caa agc att gcc ctt tta gct tca ctt ttt gag gag aac ctc 960 Thr Ile Gln Ser Ile Ala Leu Leu Ala Ser Leu Phe Glu Glu Asn Leu 305 310 315 320 att ccg cat ttg gta att gct cct cta tcg act ctg cgt aac tgg gag 1008 Ile Pro His Leu Val Ile Ala Pro Leu Ser Thr Leu Arg Asn Trp Glu 325 330 335 aga gag ttt gcc aca tgg gcc cca cag atg aac gtg gtt atg tat ttt 1056 Arg Glu Phe Ala Thr Trp Ala Pro Gln Met Asn Val Val Met Tyr Phe 340 345 350 ggc act gcg caa gct cga gca gtt atc aga gaa cat gag ttt tac tta 1104 Gly Thr Ala Gln Ala Arg Ala Val Ile Arg Glu His Glu Phe Tyr Leu 355 360 365 tcg aaa gat caa aaa aag atc aag aaa aag aaa tct gga caa ata agt 1152 Ser Lys Asp Gln Lys Lys Ile Lys Lys Lys Lys Ser Gly Gln Ile Ser 370 375 380 agc gaa agc aag caa aaa aga atc aag ttt gat gtc ctc ctc aca tcg 1200 Ser Glu Ser Lys Gln Lys Arg Ile Lys Phe Asp Val Leu Leu Thr Ser 385 390 395 400 tat gag atg atc aac cta gat tca gca gtt cta aaa cca att aag tgg 1248 Tyr Glu Met Ile Asn Leu Asp Ser Ala Val Leu Lys Pro Ile Lys Trp 405 410 415 gag tgc atg att gtt gat gaa ggt cat cga ctg aaa aat aag gat tca 1296 Glu Cys Met Ile Val Asp Glu Gly His Arg Leu Lys Asn Lys Asp Ser 420 425 430 aag ctg ttc tct tca ttg aca cag tat tca agt aac cac cgt att ctt 1344 Lys Leu Phe Ser Ser Leu Thr Gln Tyr Ser Ser Asn His Arg Ile Leu 435 440 445 ctg aca gga aca cca ctt cag aac aac ttg gat gaa ctt ttc atg ctc 1392 Leu Thr Gly Thr Pro Leu Gln Asn Asn Leu Asp Glu Leu Phe Met Leu 450 455 460 atg cat ttt ctt gat gcg ggg aag ttt gga agt ttg gag gag ttc cag 1440 Met His Phe Leu Asp Ala Gly Lys Phe Gly Ser Leu Glu Glu Phe Gln 465 470 475 480 gag gag ttc aaa gat att aat caa gag gag cag atc tca agg ttg cac 1488 Glu Glu Phe Lys Asp Ile Asn Gln Glu Glu Gln Ile Ser Arg Leu His 485 490 495 aaa atg ttg gct cca cat ttg ctc aga agg gta aaa aaa gac gta atg 1536 Lys Met Leu Ala Pro His Leu Leu Arg Arg Val Lys Lys Asp Val Met 500 505 510 aaa gac atg ccc ccc aaa aag gag ctc att ttg cgt gtt gat ctg agc 1584 Lys Asp Met Pro Pro Lys Lys Glu Leu Ile Leu Arg Val Asp Leu Ser 515 520 525 agt ctg cag aaa gaa tat tac aaa gct att ttt acc cgt aat tat caa 1632 Ser Leu Gln Lys Glu Tyr Tyr Lys Ala Ile Phe Thr Arg Asn Tyr Gln 530 535 540 gta ttg aca aaa aag gga ggt gct caa att tcc ctt aat aac att atg 1680 Val Leu Thr Lys Lys Gly Gly Ala Gln Ile Ser Leu Asn Asn Ile Met 545 550 555 560 atg gaa tta cga aaa gta tgc tgc cat cct tat atg cta gag ggt gtt 1728 Met Glu Leu Arg Lys Val Cys Cys His Pro Tyr Met Leu Glu Gly Val 565 570 575 gag cca gtt att cac gac gca aat gaa gct ttc aaa caa ctt ttg gag 1776 Glu Pro Val Ile His Asp Ala Asn Glu Ala Phe Lys Gln Leu Leu Glu 580 585 590 tct tgt gga aag ctg caa ctt cta gat aaa atg atg gtc aaa ctg aaa 1824 Ser Cys Gly Lys Leu Gln Leu Leu Asp Lys Met Met Val Lys Leu Lys 595 600 605 gag caa gga cac aga gtc cta ata tac aca cag ttt cag cat atg ctg 1872 Glu Gln Gly His Arg Val Leu Ile Tyr Thr Gln Phe Gln His Met Leu 610 615 620 gac tta ctt gaa gac tac tgt acc cat aag aaa tgg cag tac gag cga 1920 Asp Leu Leu Glu Asp Tyr Cys Thr His Lys Lys Trp Gln Tyr Glu Arg 625 630 635 640 att gat gga aag gtt ggc gga gct gag cgg caa ata cgc ata gat cgg 1968 Ile Asp Gly Lys Val Gly Gly Ala Glu Arg Gln Ile Arg Ile Asp Arg 645 650 655 ttc aat gcc aaa aat tct aac aag ttt tgt ttt ttg ctc tcc aca aga 2016 Phe Asn Ala Lys Asn Ser Asn Lys Phe Cys Phe Leu Leu Ser Thr Arg 660 665 670 gct ggt ggc tta gga ata aat ctt gca acg gct gat aca gta atc att 2064 Ala Gly Gly Leu Gly Ile Asn Leu Ala Thr Ala Asp Thr Val Ile Ile 675 680 685 tat gac agt gac tgg aat cct cat gct gat ctt caa gca atg gct aga 2112 Tyr Asp Ser Asp Trp Asn Pro His Ala Asp Leu Gln Ala Met Ala Arg 690 695 700 gct cat cga ctt ggc caa aca aat aag gtg atg att tat agg ctc ata 2160 Ala His Arg Leu Gly Gln Thr Asn Lys Val Met Ile Tyr Arg Leu Ile 705 710 715 720 aac cga ggc acc att gaa gaa agg atg atg caa ttg act aaa aag aaa 2208 Asn Arg Gly Thr Ile Glu Glu Arg Met Met Gln Leu Thr Lys Lys Lys 725 730 735 atg gtt cta gag cat ctt gtt gtt ggg aaa ctc aaa aca caa aac att 2256 Met Val Leu Glu His Leu Val Val Gly Lys Leu Lys Thr Gln Asn Ile 740 745 750 aat cag gaa gag tta gat gac atc atc agg tat gga tca aag gag ctt 2304 Asn Gln Glu Glu Leu Asp Asp Ile Ile Arg Tyr Gly Ser Lys Glu Leu 755 760 765 ttt gct agt gaa gat gat gaa gca gga aag tct gga aaa att cat tat 2352 Phe Ala Ser Glu Asp Asp Glu Ala Gly Lys Ser Gly Lys Ile His Tyr 770 775 780 gat gat gcg gct ata gac aaa ttg ctt gat cgt gat ctc gtg gag gca 2400 Asp Asp Ala Ala Ile Asp Lys Leu Leu Asp Arg Asp Leu Val Glu Ala 785 790 795 800 gag gaa gtc tca gtg gat gat gaa gag gag aat gga ttc tta aag gct 2448 Glu Glu Val Ser Val Asp Asp Glu Glu Glu Asn Gly Phe Leu Lys Ala 805 810 815 ttc aag gtg gct aat ttt gaa tat ata gat gaa aat gag gca gca gca 2496 Phe Lys Val Ala Asn Phe Glu Tyr Ile Asp Glu Asn Glu Ala Ala Ala 820 825 830 tta gag gca cag aga gtc gct gct gaa agc aaa tct tca gca ggc aat 2544 Leu Glu Ala Gln Arg Val Ala Ala Glu Ser Lys Ser Ser Ala Gly Asn 835 840 845 tct gat aga gca agt tat tgg gaa gag ttg tta aaa gat aaa ttt gag 2592 Ser Asp Arg Ala Ser Tyr Trp Glu Glu Leu Leu Lys Asp Lys Phe Glu 850 855 860 ctg cac cag gct gag gag ctt aat gct ctt gga aaa agg aag aga agt 2640 Leu His Gln Ala Glu Glu Leu Asn Ala Leu Gly Lys Arg Lys Arg Ser 865 870 875 880 cgc aag cag ttg gta tcc att gaa gaa gat gat ctt gct ggt ttg gaa 2688 Arg Lys Gln Leu Val Ser Ile Glu Glu Asp Asp Leu Ala Gly Leu Glu 885 890 895 gat gtg agc tct gat gga gat gaa agt tat gaa gct gag tca aca gat 2736 Asp Val Ser Ser Asp Gly Asp Glu Ser Tyr Glu Ala Glu Ser Thr Asp 900 905 910 ggt gaa gca gca gga caa gga gtt cag acg ggt cga cgg ccg tac aga 2784 Gly Glu Ala Ala Gly Gln Gly Val Gln Thr Gly Arg Arg Pro Tyr Arg 915 920 925 aga aag ggt cgc gat aat ttg gaa cca act ccg ttg atg gaa ggt gag 2832 Arg Lys Gly Arg Asp Asn Leu Glu Pro Thr Pro Leu Met Glu Gly Glu 930 935 940 ggg aga tct ttc aga gta ctg ggt ttc aac cag agt caa agg gcc att 2880 Gly Arg Ser Phe Arg Val Leu Gly Phe Asn Gln Ser Gln Arg Ala Ile 945 950 955 960 ttt gta cag act ttg atg agg tat gga gct ggc aat ttt gat tgg aag 2928 Phe Val Gln Thr Leu Met Arg Tyr Gly Ala Gly Asn Phe Asp Trp Lys 965 970 975 gag ttt gtt cct cgc tta aag cag aag acc ttt gaa gaa ata aat gaa 2976 Glu Phe Val Pro Arg Leu Lys Gln Lys Thr Phe Glu Glu Ile Asn Glu 980 985 990 tat gga ata ctc ttc ttg aag cac att gct gaa gaa ata gac gag aat 3024 Tyr Gly Ile Leu Phe Leu Lys His Ile Ala Glu Glu Ile Asp Glu Asn 995 1000 1005 tct cca acc ttt tca gat ggt gtg ccc aag gaa gga ctt aga ata 3069 Ser Pro Thr Phe Ser Asp Gly Val Pro Lys Glu Gly Leu Arg Ile 1010 1015 1020 gaa gat gtt cta gtc aga att gct ctt ctg ata cta gtt cag gag 3114 Glu Asp Val Leu Val Arg Ile Ala Leu Leu Ile Leu Val Gln Glu 1025 1030 1035 aag gtg aaa ttt gta gaa gat cat cca ggg aaa cct gtt ttc ccc 3159 Lys Val Lys Phe Val Glu Asp His Pro Gly Lys Pro Val Phe Pro 1040 1045 1050 tct cgc att ctt gaa aga ttc ccc gga ctg aga agt gga aaa att 3204 Ser Arg Ile Leu Glu Arg Phe Pro Gly Leu Arg Ser Gly Lys Ile 1055 1060 1065 tgg aag gag gaa cat gac aag ata atg ata cgt gct gtt tta aag 3249 Trp Lys Glu Glu His Asp Lys Ile Met Ile Arg Ala Val Leu Lys 1070 1075 1080 cat ggg tac gga cgg tgg caa gct att gtt gat gac aaa gag ttg 3294 His Gly Tyr Gly Arg Trp Gln Ala Ile Val Asp Asp Lys Glu Leu 1085 1090 1095 ggg atc caa gag ctt atc tgc aaa gaa ttg aat ttc cct cac ata 3339 Gly Ile Gln Glu Leu Ile Cys Lys Glu Leu Asn Phe Pro His Ile 1100 1105 1110 agt ttg tct gct gct gaa caa gct ggt ttg cag ggg cag aat ggt 3384 Ser Leu Ser Ala Ala Glu Gln Ala Gly Leu Gln Gly Gln Asn Gly 1115 1120 1125 agt ggg ggc tct aat ccg gga gca cag act aac cag aat cct gga 3429 Ser Gly Gly Ser Asn Pro Gly Ala Gln Thr Asn Gln Asn Pro Gly 1130 1135 1140 agc gtt att act ggg aac aat aat gct tct gct gat ggg gct caa 3474 Ser Val Ile Thr Gly Asn Asn Asn Ala Ser Ala Asp Gly Ala Gln 1145 1150 1155 gta aac tcg atg ttc tat tat cgg gac atg cag aga cga ctt gtt 3519 Val Asn Ser Met Phe Tyr Tyr Arg Asp Met Gln Arg Arg Leu Val 1160 1165 1170 gag ttt gtg aaa aag cga gtt ctg ctt ttg gag aag gcg atg aat 3564 Glu Phe Val Lys Lys Arg Val Leu Leu Leu Glu Lys Ala Met Asn 1175 1180 1185 tat gaa tac gca gag gaa tat tat gga ctt ggt ggc tca tca tct 3609 Tyr Glu Tyr Ala Glu Glu Tyr Tyr Gly Leu Gly Gly Ser Ser Ser 1190 1195 1200 atc cct act gaa gaa cca gaa gct gaa cca aag atc gct gac aca 3654 Ile Pro Thr Glu Glu Pro Glu Ala Glu Pro Lys Ile Ala Asp Thr 1205 1210 1215 gtg gga gtg agc ttt att gag gtt gat gat gaa atg ctt gat gga 3699 Val Gly Val Ser Phe Ile Glu Val Asp Asp Glu Met Leu Asp Gly 1220 1225 1230 ctt cct aag act gat cct atc act tca gaa gaa att atg ggg gct 3744 Leu Pro Lys Thr Asp Pro Ile Thr Ser Glu Glu Ile Met Gly Ala 1235 1240 1245 gct gtt gac aac aac caa gcg cgg gtc gaa ata gct caa cat tat 3789 Ala Val Asp Asn Asn Gln Ala Arg Val Glu Ile Ala Gln His Tyr 1250 1255 1260 aac cag atg tgc aaa ctt ctt gat gag aac gct cgg gaa tca gtc 3834 Asn Gln Met Cys Lys Leu Leu Asp Glu Asn Ala Arg Glu Ser Val 1265 1270 1275 caa gca tat gta aac aac caa cca ccg agt acc aag gtg aat gag 3879 Gln Ala Tyr Val Asn Asn Gln Pro Pro Ser Thr Lys Val Asn Glu 1280 1285 1290 agc ttc cgt gca ctc aaa tct atc aat ggt aac att aac aca atc 3924 Ser Phe Arg Ala Leu Lys Ser Ile Asn Gly Asn Ile Asn Thr Ile 1295 1300 1305 ctt tcg att aca tct gat caa tcc aag tca cat gaa gac gac acc 3969 Leu Ser Ile Thr Ser Asp Gln Ser Lys Ser His Glu Asp Asp Thr 1310 1315 1320 aag cca gac cta aac aat gtt gag atg aag gac acg gcc gaa gaa 4014 Lys Pro Asp Leu Asn Asn Val Glu Met Lys Asp Thr Ala Glu Glu 1325 1330 1335 aca aaa ccg tta aga ggt ggc gtc gtc gat ctg aat gtg gtg gag 4059 Thr Lys Pro Leu Arg Gly Gly Val Val Asp Leu Asn Val Val Glu 1340 1345 1350 gga gag gag aac att gct gaa gct agt gga agt gtt gat gta aaa 4104 Gly Glu Glu Asn Ile Ala Glu Ala Ser Gly Ser Val Asp Val Lys 1355 1360 1365 atg gaa gaa gcc aaa gaa gaa gag aag cca aag aac atg gtc gtt 4149 Met Glu Glu Ala Lys Glu Glu Glu Lys Pro Lys Asn Met Val Val 1370 1375 1380 gat tga 4155 Asp 2 1384 PRT Arabidopsis thaliana 2 Met Ser Ser Leu Val Glu Arg Leu Arg Ile Arg Ser Asp Arg Lys Pro 1 5 10 15 Val Tyr Asn Leu Asp Asp Ser Asp Asp Asp Asp Phe Val Pro Lys Lys 20 25 30 Asp Arg Thr Phe Glu Gln Val Glu Ala Ile Val Arg Thr Asp Ala Lys 35 40 45 Glu Asn Ala Cys Gln Ala Cys Gly Glu Ser Thr Asn Leu Val Ser Cys 50 55 60 Asn Thr Cys Thr Tyr Ala Phe His Ala Lys Cys Leu Val Pro Pro Leu 65 70 75 80 Lys Asp Ala Ser Val Glu Asn Trp Arg Cys Pro Glu Cys Val Ser Pro 85 90 95 Leu Asn Glu Ile Asp Lys Ile Leu Asp Cys Glu Met Arg Pro Thr Lys 100 105 110 Ser Ser Glu Gln Gly Ser Ser Asp Ala Glu Pro Lys Pro Ile Phe Val 115 120 125 Lys Gln Tyr Leu Val Lys Trp Lys Gly Leu Ser Tyr Leu His Cys Ser 130 135 140 Trp Val Pro Glu Lys Glu Phe Gln Lys Ala Tyr Lys Ser Asn His Arg 145 150 155 160 Leu Lys Thr Arg Val Asn Asn Phe His Arg Gln Met Glu Ser Phe Asn 165 170 175 Asn Ser Glu Asp Asp Phe Val Ala Ile Arg Pro Glu Trp Thr Thr Val 180 185 190 Asp Arg Ile Leu Ala Cys Arg Glu Glu Asp Gly Glu Leu Glu Tyr Leu 195 200 205 Val Lys Tyr Lys Glu Leu Ser Tyr Asp Glu Cys Tyr Trp Glu Ser Glu 210 215 220 Ser Asp Ile Ser Thr Phe Gln Asn Glu Ile Gln Arg Phe Lys Asp Val 225 230 235 240 Asn Ser Arg Thr Arg Arg Ser Lys Asp Val Asp His Lys Arg Asn Pro 245 250 255 Arg Asp Phe Gln Gln Phe Asp His Thr Pro Glu Phe Leu Lys Gly Leu 260 265 270 Leu His Pro Tyr Gln Leu Glu Gly Leu Asn Phe Leu Arg Phe Ser Trp 275 280 285 Ser Lys Gln Thr His Val Ile Leu Ala Asp Glu Met Gly Leu Gly Lys 290 295 300 Thr Ile Gln Ser Ile Ala Leu Leu Ala Ser Leu Phe Glu Glu Asn Leu 305 310 315 320 Ile Pro His Leu Val Ile Ala Pro Leu Ser Thr Leu Arg Asn Trp Glu 325 330 335 Arg Glu Phe Ala Thr Trp Ala Pro Gln Met Asn Val Val Met Tyr Phe 340 345 350 Gly Thr Ala Gln Ala Arg Ala Val Ile Arg Glu His Glu Phe Tyr Leu 355 360 365 Ser Lys Asp Gln Lys Lys Ile Lys Lys Lys Lys Ser Gly Gln Ile Ser 370 375 380 Ser Glu Ser Lys Gln Lys Arg Ile Lys Phe Asp Val Leu Leu Thr Ser 385 390 395 400 Tyr Glu Met Ile Asn Leu Asp Ser Ala Val Leu Lys Pro Ile Lys Trp 405 410 415 Glu Cys Met Ile Val Asp Glu Gly His Arg Leu Lys Asn Lys Asp Ser 420 425 430 Lys Leu Phe Ser Ser Leu Thr Gln Tyr Ser Ser Asn His Arg Ile Leu 435 440 445 Leu Thr Gly Thr Pro Leu Gln Asn Asn Leu Asp Glu Leu Phe Met Leu 450 455 460 Met His Phe Leu Asp Ala Gly Lys Phe Gly Ser Leu Glu Glu Phe Gln 465 470 475 480 Glu Glu Phe Lys Asp Ile Asn Gln Glu Glu Gln Ile Ser Arg Leu His 485 490 495 Lys Met Leu Ala Pro His Leu Leu Arg Arg Val Lys Lys Asp Val Met 500 505 510 Lys Asp Met Pro Pro Lys Lys Glu Leu Ile Leu Arg Val Asp Leu Ser 515 520 525 Ser Leu Gln Lys Glu Tyr Tyr Lys Ala Ile Phe Thr Arg Asn Tyr Gln 530 535 540 Val Leu Thr Lys Lys Gly Gly Ala Gln Ile Ser Leu Asn Asn Ile Met 545 550 555 560 Met Glu Leu Arg Lys Val Cys Cys His Pro Tyr Met Leu Glu Gly Val 565 570 575 Glu Pro Val Ile His Asp Ala Asn Glu Ala Phe Lys Gln Leu Leu Glu 580 585 590 Ser Cys Gly Lys Leu Gln Leu Leu Asp Lys Met Met Val Lys Leu Lys 595 600 605 Glu Gln Gly His Arg Val Leu Ile Tyr Thr Gln Phe Gln His Met Leu 610 615 620 Asp Leu Leu Glu Asp Tyr Cys Thr His Lys Lys Trp Gln Tyr Glu Arg 625 630 635 640 Ile Asp Gly Lys Val Gly Gly Ala Glu Arg Gln Ile Arg Ile Asp Arg 645 650 655 Phe Asn Ala Lys Asn Ser Asn Lys Phe Cys Phe Leu Leu Ser Thr Arg 660 665 670 Ala Gly Gly Leu Gly Ile Asn Leu Ala Thr Ala Asp Thr Val Ile Ile 675 680 685 Tyr Asp Ser Asp Trp Asn Pro His Ala Asp Leu Gln Ala Met Ala Arg 690 695 700 Ala His Arg Leu Gly Gln Thr Asn Lys Val Met Ile Tyr Arg Leu Ile 705 710 715 720 Asn Arg Gly Thr Ile Glu Glu Arg Met Met Gln Leu Thr Lys Lys Lys 725 730 735 Met Val Leu Glu His Leu Val Val Gly Lys Leu Lys Thr Gln Asn Ile 740 745 750 Asn Gln Glu Glu Leu Asp Asp Ile Ile Arg Tyr Gly Ser Lys Glu Leu 755 760 765 Phe Ala Ser Glu Asp Asp Glu Ala Gly Lys Ser Gly Lys Ile His Tyr 770 775 780 Asp Asp Ala Ala Ile Asp Lys Leu Leu Asp Arg Asp Leu Val Glu Ala 785 790 795 800 Glu Glu Val Ser Val Asp Asp Glu Glu Glu Asn Gly Phe Leu Lys Ala 805 810 815 Phe Lys Val Ala Asn Phe Glu Tyr Ile Asp Glu Asn Glu Ala Ala Ala 820 825 830 Leu Glu Ala Gln Arg Val Ala Ala Glu Ser Lys Ser Ser Ala Gly Asn 835 840 845 Ser Asp Arg Ala Ser Tyr Trp Glu Glu Leu Leu Lys Asp Lys Phe Glu 850 855 860 Leu His Gln Ala Glu Glu Leu Asn Ala Leu Gly Lys Arg Lys Arg Ser 865 870 875 880 Arg Lys Gln Leu Val Ser Ile Glu Glu Asp Asp Leu Ala Gly Leu Glu 885 890 895 Asp Val Ser Ser Asp Gly Asp Glu Ser Tyr Glu Ala Glu Ser Thr Asp 900 905 910 Gly Glu Ala Ala Gly Gln Gly Val Gln Thr Gly Arg Arg Pro Tyr Arg 915 920 925 Arg Lys Gly Arg Asp Asn Leu Glu Pro Thr Pro Leu Met Glu Gly Glu 930 935 940 Gly Arg Ser Phe Arg Val Leu Gly Phe Asn Gln Ser Gln Arg Ala Ile 945 950 955 960 Phe Val Gln Thr Leu Met Arg Tyr Gly Ala Gly Asn Phe Asp Trp Lys 965 970 975 Glu Phe Val Pro Arg Leu Lys Gln Lys Thr Phe Glu Glu Ile Asn Glu 980 985 990 Tyr Gly Ile Leu Phe Leu Lys His Ile Ala Glu Glu Ile Asp Glu Asn 995 1000 1005 Ser Pro Thr Phe Ser Asp Gly Val Pro Lys Glu Gly Leu Arg Ile 1010 1015 1020 Glu Asp Val Leu Val Arg Ile Ala Leu Leu Ile Leu Val Gln Glu 1025 1030 1035 Lys Val Lys Phe Val Glu Asp His Pro Gly Lys Pro Val Phe Pro 1040 1045 1050 Ser Arg Ile Leu Glu Arg Phe Pro Gly Leu Arg Ser Gly Lys Ile 1055 1060 1065 Trp Lys Glu Glu His Asp Lys Ile Met Ile Arg Ala Val Leu Lys 1070 1075 1080 His Gly Tyr Gly Arg Trp Gln Ala Ile Val Asp Asp Lys Glu Leu 1085 1090 1095 Gly Ile Gln Glu Leu Ile Cys Lys Glu Leu Asn Phe Pro His Ile 1100 1105 1110 Ser Leu Ser Ala Ala Glu Gln Ala Gly Leu Gln Gly Gln Asn Gly 1115 1120 1125 Ser Gly Gly Ser Asn Pro Gly Ala Gln Thr Asn Gln Asn Pro Gly 1130 1135 1140 Ser Val Ile Thr Gly Asn Asn Asn Ala Ser Ala Asp Gly Ala Gln 1145 1150 1155 Val Asn Ser Met Phe Tyr Tyr Arg Asp Met Gln Arg Arg Leu Val 1160 1165 1170 Glu Phe Val Lys Lys Arg Val Leu Leu Leu Glu Lys Ala Met Asn 1175 1180 1185 Tyr Glu Tyr Ala Glu Glu Tyr Tyr Gly Leu Gly Gly Ser Ser Ser 1190 1195 1200 Ile Pro Thr Glu Glu Pro Glu Ala Glu Pro Lys Ile Ala Asp Thr 1205 1210 1215 Val Gly Val Ser Phe Ile Glu Val Asp Asp Glu Met Leu Asp Gly 1220 1225 1230 Leu Pro Lys Thr Asp Pro Ile Thr Ser Glu Glu Ile Met Gly Ala 1235 1240 1245 Ala Val Asp Asn Asn Gln Ala Arg Val Glu Ile Ala Gln His Tyr 1250 1255 1260 Asn Gln Met Cys Lys Leu Leu Asp Glu Asn Ala Arg Glu Ser Val 1265 1270 1275 Gln Ala Tyr Val Asn Asn Gln Pro Pro Ser Thr Lys Val Asn Glu 1280 1285 1290 Ser Phe Arg Ala Leu Lys Ser Ile Asn Gly Asn Ile Asn Thr Ile 1295 1300 1305 Leu Ser Ile Thr Ser Asp Gln Ser Lys Ser His Glu Asp Asp Thr 1310 1315 1320 Lys Pro Asp Leu Asn Asn Val Glu Met Lys Asp Thr Ala Glu Glu 1325 1330 1335 Thr Lys Pro Leu Arg Gly Gly Val Val Asp Leu Asn Val Val Glu 1340 1345 1350 Gly Glu Glu Asn Ile Ala Glu Ala Ser Gly Ser Val Asp Val Lys 1355 1360 1365 Met Glu Glu Ala Lys Glu Glu Glu Lys Pro Lys Asn Met Val Val 1370 1375 1380 Asp 3 9353 DNA Arabidopsis thaliana 3 atgagtagtt tggtggagag gcttcgcata cgatctgata ggaaaccagt ttataaccta 60 gatgattctg atgatgacga cttcgttcct aaaaaagatc gaacctttga gcaagtcgag 120 gctattgtca gaactgatgc ggtttgtttc tcctctcgag cttattgttc agcttttact 180 gttttatgtg ttctatttta atcctttttt ttgtgttgtt actctgaatt tgtagaaaga 240 aaatgcatgt caggcttgtg gggaaagtac taatcttgta agctgcaata catgcactta 300 tgcgttccat gctaaatgct tagttccacc tcttaaagat gcttccgtgg aaaattggag 360 atgccctgaa tgtgtaagat tttagttacg gtccacaatt atgttttggg atgctacagg 420 ttccattttt cttacatgga agaattgttg tttacatttg caggttagtc ctcttaacga 480 gatagataag atattggatt gtgaaatgcg tcctacaaaa tctagtgaac aaggttcctc 540 cgatgcggaa ccgaagccaa tttttgtgaa acagtatctc gtgaagtgga agggattatc 600 ataccttcac tgctcttggt agttactgcg tgtctttttt gctgtctgga cacgctaatt 660 atcaatgttt ctttctgtga acactataat atgtgattta tttcctttta ctaatcatag 720 ggtgcctgag aaggagttcc agaaggctta taagtcaaat catcgtttaa aaaccagagt 780 gaacaatttt caccgtcaaa tggagtcatt caataacagc gaagatgatt ttgttgccat 840 acgtcctgag tggaccactg ttgatcggat tcttgcctgc aggtctagag aatggaatta 900 attcctttat ttatctatct gccaactttt tttttaatat ccttgttttc agcataatcc 960 attctctaat aaacacgtat ctttgataga gtgctgctta acctaaattt actgttatca 1020 cgattttggg tctctgaaac atgataaatg acctgcttac cttttttttc ttctttttaa 1080 gttaccattt tcttagttgt ttcgtaaatc aggaattgtg acagttgcat tggtttcttt 1140 tatgatatag agaggaagat ggggagctgg aatatcttgt caaatataaa gagctatcct 1200 atgatgaatg ttattgggag tcagaatcag acatctcaac cttccagaat gaaattcaaa 1260 ggttcaagga tgtaaattct agaactcgca gaagtaaaga tgttgaccat aaaagaaatc 1320 ccagagactt tcaacagttt gatcatactc ctgaattcct caaaggtatt tggatcacct 1380 taaatcatat actataaatg tttcttatat ttggtactta tagatgttat gatttatttg 1440 tttcctgcga ttgaaggctt gttacatcca taccagcttg agggacttaa ttttttgcgg 1500 ttctcgtggt caaaacagac gcatgtaatc cttgctgatg aaatgggact aggtaatttt 1560 tcaattgtcc cacttgggtg gtcacataga tcttttcatc cattgtaagg ggcctttgtt 1620 ttctattcct gtaatgttgt gagatttttc ctgttacagg caagacaatt caaagcattg 1680 cccttttagc ttcacttttt gaggagaacc tcattccgca tttggtaatt gctcctctat 1740 cgactctgcg taactgggag agagagtttg ccacatgggc cccacagatg aacgtggtat 1800 gtatgcagtt atacacgcaa tgatctgtgc catttgtatg tttttgttgt ttgttaatgg 1860 aatggtcttc gtggtcattt gacgggtagg ttatgtattt tggcactgcg caagctcgag 1920 cagttatcag agaacatgag ttttacttat cgaaagatca aaaaaagatc aagaaaaaga 1980 aatctggaca aataagtagc gaaagcaagc aaaaaagaat caagtttgat gtcctcctca 2040 catcgtatga gatgatcaac ctagattcag cagttctaaa accaattaag tgggagtgca 2100 tggtaactct tattctctaa tgagacttta ctttctctta gtcgtctctc tttctctctt 2160 acatgttgcc tagtaacaat tgttttgggc agattgttga tgaaggtcat cgactgaaaa 2220 ataaggattc aaagctgttc tcttcattga cacagtattc aagtaaccac cgtattcttc 2280 tgacaggaac accacttcag gttcgtcatt tgagtttgat ttctgaagtt tatactttca 2340 atagttgtat ctgagcatag tagctacgat ttgcaatgag aattgttata tattatcttg 2400 cactaatgtc ttacctgatt agttgcaata tgttactgat gattatgtgg tgcctttaca 2460 gaacaacttg gatgaacttt tcatgctcat gcattttctt gatgcgggga aggtatcaca 2520 agaatagcaa agataaataa gttcgcatac ttaacagaat tttatgtagc taacatgtta 2580 tttgattgca caatacttgc agtttggaag tttggaggag ttccaggagg agttcaaaga 2640 tattaatcaa gaggagcaga tctcaaggtt gcacaaaatg ttggctccac atttgctcag 2700 aagtattaac caaaactatt tgttcatctt ttttaattta tatgtgtttc aaaagtttgg 2760 ttggagggaa tctttcatag taataatttt atgatcttaa ccatgctgtc tcgtattttg 2820 attgctcttc caggggtaaa aaaagacgta atgaaagaca tgccccccaa aaaggagctc 2880 attttgcgtg ttgatctgag cagtctgcag aaagaatatt acaaagctat ttttacccgt 2940 aattatcaag tattgacaaa aaagggaggt gctcaagtaa gttcttttta atttttgttt 3000 acactttttg gatcattaaa cctcataggt ggggtagaaa ccaggtcaac tgtaatcgtc 3060 tagtgaatgt attggtctat ttctgtttca gatttccctt aataacatta tgatggaatt 3120 acgaaaagta tgctgccatc cttatatgct agagggtgtt gagccagtta ttcacgacgc 3180 aaatgaagct ttcaagtaat atctcatttc ccaaaaatgg ttatctgttt attactactt 3240 attaaagtcg tctgctaact tttgcgttga acgttttctt atatgtatca aagacaactt 3300 ttggagtctt gtggaaagct gcaacttcta gataaaatga tggtcaaact gaaagagcaa 3360 ggacacagag tcctaatata cacacagttt cagcatatgc tggacttact tgaagactac 3420 tgtacccata aggtatttga acttcttata tgtacagtct gtttcagtag attttcattc 3480 ttgttgtttt tgtagaatat cattttgaca ctgtagaatc aactctacca ttttctagtg 3540 ttagagtact taggcacaat tatggaaata caagcatgtg ctgaaattga gagtatatga 3600 gcattctgtg cccaactgaa agagcaaaga cacaaagttt ccttataaac acagtacaaa 3660 tcacaagttt agccatcttc tatgtacagt agttttccaa taggtcgagc atgtgctgaa 3720 actgtgtgta cagagttctc ataaacacac agtttcagca tatgctggat ctacttgaag 3780 actactgttc ttataaggta ctgaacttgt tatctgtact gcgtatatac gagatctctg 3840 tattcttgct cttttatttt gacactttgt tctcatatac actcggttca gcacatgctc 3900 gacttactgc ctaaggatct tgaaaaaggt agagttgatt ctatgtctag gtgcaattac 3960 tttcttagaa tttttgtcat tacttactct gttggcaata taacttcttt attccctcaa 4020 agattacttt ttttggtttc ttgaaatgcc attatcaata ccattgcttt tgctgacgca 4080 tgcacttgag acaacttgtt tttatctctt tctagcacat ttttttttaa catgcagtta 4140 aggaaaattc tcatatgatt tacgctgttc attttcttgt ctttgtcaga aatggcagta 4200 cgagcgaatt gatggaaagg ttggcggagc tgagcggcaa atacgcatag atcggttcaa 4260 tgccaaaaat tctaacaagt tttgtttttt gctctccaca agagctggtg gcttaggaat 4320 aaatcttgca acggctgata cagtaatcat ttatgacagg tttgaatttc agcttctctt 4380 agtgtcatct gtactctttt catagttatt gtgtcaagct gtaagaggaa ctatttggct 4440 tgatagcata atattttgga agtttaatgt tgatttttaa gtgaattggg ttgtgatgag 4500 tgataaaaag gcacttggct tttttccaat aacagctatt tcttgaacat ggatgttcta 4560 agacagcagg aagatcagga aaattattaa ccgctatctt gctaataatt agattttgta 4620 ggcatgcaat atgggtggcg tccatgggat cctgcttgga tggcagtttg ttttggttta 4680 cgcctgttca cattttcata cgtacgattg aaactgtttt atctgtttct gtagtgactg 4740 gaatcctcat gctgatcttc aagcaatggc tagagctcat cgacttggcc aaacaaataa 4800 ggttttaaat tttatctctt agtgctgtca acttgcaatt ttgtgttctt ttttgtagtt 4860 tccctaattt tccttatatt ttcctttagg tgatgattta taggctcata aaccgaggca 4920 ccattgaaga aaggatgatg caattgacta aaaagaaaat ggttctagag catcttgttg 4980 ttgggaaact caaaacacaa aacattaatc aggtaaactt ttattgcttg aagccttttt 5040 acttgattac aaatttctca acggattgga gctggaaggt agaaattcca agaagaacac 5100 cttcggttat aacttataag tgtgaaatta aaagataaaa actttagaga gaaggggtcc 5160 atatttgtta attgtttgtc actaagtatg tgtttgtttt gttttcctga ctgcaattta 5220 ggaagagtta gatgacatca tcaggtatgg atcaaaggag ctttttgcta gtgaagatga 5280 tgaagcagga aagtctggaa aaattcatta tgatgatgcg gctatagaca agtaatagac 5340 tccttactct tttcctcttg ttttgttttt gattaacaag gatatctgat ctttccgatt 5400 gctcctttct tatgaaagct tttgcagtca attgcatggg cgtatttcat tatttgtctc 5460 tatcttctgt tctgcagatt gcttgatcgt gatctcgtgg aggcagagga agtctcagtg 5520 gatgatgaag aggagaatgg attcttaaag gctttcaagg ttttcttgcc tcttactatt 5580 cttcctcttc tattagtttt ctctgaatca gtgtttactg atttcaatgc tccattggag 5640 tctatgctta attgtattct tatattccat gatattcaga ctgtggttgg ctatcgaaat 5700 cccttctgct gtgcacaatc ttgtcaaatc attacgtgct aagtttgtag gatcaataca 5760 ctttatgcca gttcgctttg atgcttatag acagtcttta gaaagtgtct attgattgtt 5820 cgttccggct caatgtgaaa gccaacttaa tgaaaattag tgatgatgac ttaagttaga 5880 aatttatgct tgtggtgatg ttgattgagc caatttattg atttggttat atttcttttg 5940 aaccctgatc atattgaatg cgttatatga gtggtcttta gacttagctg gaacataagg 6000 ctgtgtcctg cattgctgct tgtcacctct taatattcga actccctaaa acattgtttg 6060 tctttgtgtg catatagaac tgttctgaag caaatagggt gtctggtact gtttagtgtc 6120 attaactctg aaaatgattt cccttgtaag attctgtgat cttcctgtat tgtaggtggc 6180 taattttgaa tatatagatg aaaatgaggc agcagcatta gaggcacaga gagtcgctgc 6240 tgaaagcaaa tcttcagcag gcaattctga tagagcaagt tattgggaag agttgttaaa 6300 agataaattt gagctgcacc aggctgagga gcttaatgct cttggaaaaa ggaagagaag 6360 tcgcaagcag gtttggtctc ttcttgatcc cccttatcca attgtggcat catattgata 6420 actggatttt tcaccattta tgttctttct gattctgtcc tgtttcatat atttattcat 6480 gttgtctaac ttttcctttt gaattcctta ggtagctaaa ttcagaaagt aataatttag 6540 ttgactgtat ccttctaaat tgagaaagta taatttagtt gactgtatcc agtataaaac 6600 taaacgccct tgtcctccta tcaactggtt tgacagatct tatgggttta catgttggat 6660 caagtaattg gggttggtag aggctcaatt aactatagtc ttctgttttc ctctgcaaga 6720 aatacgtttt gtttcactct ctaacttgat atagctcaat tactgacaat atacattggt 6780 ttggtctgcc atcatcgttt catgtctttc aataaaggct gttctaattc ttctatggga 6840 tttttttcat agttggtatc cattgaagaa gatgatcttg ctggtttgga agatgtgagc 6900 tctgatggag atgaaagtta tgaagctgag tcaacagatg gtgaagcagc aggacaagga 6960 gttcagacgg gtcgacggcc gtacagaaga aagggtcgcg gtattaccac gtttcggatt 7020 taatttaatt tgtaatggag ctgaaaatga ctgatattag aagtgtgcgc agtttattag 7080 atgagttttt tttctataga taatttggaa ccaactccgt tgatggaagg tgaggggaga 7140 tctttcagag tactgggttt caaccagagt caaagggcca tttttgtaca gactttgatg 7200 aggtatctac tttccattaa ggcctttaga cgccagaagc tattctgtct aaattttaca 7260 gtttcatccc ccgatgcatc taaattatca tcagtcttgt ggtgctcaat atttacaagt 7320 ttttccggtt ggacaaaata attgcaggta tggagctggc aattttgatt ggaaggagtt 7380 tgttcctcgc ttaaagcaga agacctttga agaaataaat gagtacgggc tcaacccttt 7440 aatgctcttc tcttctgctt ctttacaaaa aacgcatcat tataaaaagg ctttctggtt 7500 tattctttaa ctaatttttt aatgactgtt tctcagatat ggaatactct tcttgaagca 7560 cattgctgaa gaaatagacg agaattctcc aaccttttca ggtgatcgat aattgatatt 7620 ttcactgttt gctgcttttc cctaaatgag atcattgctt ctcctgttaa ccggttaaat 7680 gtattaatat aatggtcgtt gtctatagat ggtgtgccca aggaaggact tagaatagaa 7740 gatgttctag tcagaattgc tcttctgata ctagttcagg agaaggtgag tctattgact 7800 ttaattcttc attaagttct ctcttttata tctgagtttt tttttggtat atgttacttc 7860 tagtctatag tttagctctg tacataagtt tttaatacag taatgtatgt tcaaacctca 7920 ctaagatttg gatcccgggt tacttatgtt tttttggtgc tctggcccga caggtgaaat 7980 ttgtagaaga tcatccaggg aaacctgttt tcccctctcg cattcttgaa agattccccg 8040 gactgagaag tggaaaaatt tggaaggagg aacatgacaa gataatgata cgtgctgttt 8100 taaagtatga accctgcacc actgttctta ccgaatggtt ttattttctc atcattctcc 8160 attacttgct cacattttct tttccttctc tggaaatttg aatctttagg catgggtacg 8220 gacggtggca agctattgtt gatgacaaag agttggggat ccaagagctt atctgcaaag 8280 aattgaattt ccctcacata agtttgtctg ctgctgaaca agctggtttg caggggcaga 8340 atggtagtgg gggctctaat ccgggagcac agactaacca gaatcctgga agcgttatta 8400 ctgggaacaa taatgcttct gctgatgggg ctcaagtaaa ctcgatgttc tattatcggg 8460 acatgcagag acgacttgtt gagtttgtga aaaagcgagt tctgcttttg gagaaggcga 8520 tgaattatga atacgcagag gaatattatg tatgttgtac catctgcagt gttggtactt 8580 actcacatgt tttgcgctga attgtttaac tttgattgaa tctctggttg cagggacttg 8640 gtggctcatc atctatccct actgaagaac cagaagctga accaaagatc gctgacacag 8700 tgggagtgag ctttattgag gttgatgatg aaatgcttga tggacttcct aagactgatc 8760 ctatcagtaa gttccatcac aagtttcttt atttaacgag ttgttgattc taatgtgagc 8820 tctctgaatc tcgctgcagc ttcagaagaa attatggggg ctgctgttga caacaaccaa 8880 gcgcgggtcg aaatagctca acattataac caggtaagct atgctttttt cctttggtgg 8940 taggctaatg tctagaacta gtatatcaca ctaatatctc tccggttatt cagatgtgca 9000 aacttcttga tgagaacgct cgggaatcag tccaagcata tgtaaacaac caaccaccga 9060 gtaccaaggt gaatgagagc ttccgtgcac tcaaatctat caatggtaac attaacacaa 9120 tcctttcgat tacatctgat caatccaagt cacatgaaga cgacaccaag ccagacctaa 9180 acaatgttga gatgaaggac acggccgaag aaacaaaacc gttaagaggt ggcgtcgtcg 9240 atctgaatgt ggtggaggga gaggagaaca ttgctgaagc tagtggaagt gttgatgtaa 9300 aaatggaaga agccaaagaa gaagagaagc caaagaacat ggtcgttgat tga 9353 4 1403 DNA Arabidopsis thaliana 4 atgaacctta aggagacgga gctttgtctt ggcctccccg gaggcactga aaccgttgaa 60 agtccggcca agtcgggtgt tgggaacaag agaggcttct ccgagaccgt tgatctcaaa 120 cttaatcttc aatctaacaa acaaggacat gtggatctca acactaatgg agctcccaag 180 gagaagacct tccttaaaga cccttctaag cctcctgcta agtaagttct atttacacaa 240 ttccttaaga agaagacctt ccttaaaagg gaagactttt tttttttttt tttgagataa 300 aaagactaat agttgatata aaagttctta aaatacatat atatgaaaga tgtaaggatg 360 cataagtaat aacgttattg aatgtgtgtg tgtgttgtta tattctatgc agagcacaag 420 tggtgggttg gccatcggtg aggaactacc ggaaaaatgt tatggctaat cagaagagcg 480 gcgaagcaga ggaggcaatg agtagtggtg gaggaaccgt cgcctttgtg aaggtttcca 540 tggatggagc tccttatctt cggaaggttg acctcaagat gtacaccagc tacaaggatc 600 tctctgatgc cttggccaaa atgttcagct cctttaccat gggtatgcat tttcagacat 660 ataagtcgaa ttatcattat tatttttgtg tttacttaca attttttctt tttaacgata 720 cagttttttc catatacgac taattaatat gataagtttt gggattttga ttaattaagg 780 gagttatgga gcacaaggga tgatagattt catgaacgag agtaaagtga tggatctgtt 840 gaacagttct gagtatgttc caagctacga ggacaaagat ggtgactgga tgctcgttgg 900 tgatgtcccc tggccgtgag tttcctcatt cttcttgctt tcattattat gaccaaaatt 960 attctctaaa caaaaaaaac aatattctct aaagcattat tattgatatt acttatcaaa 1020 aaaatacaca aaatgataat caatatccat gtgttataaa cacgcacagc catcttttgg 1080 ttggcatggg acagaactca gagacagaga agatgtttat atataaatac taactcatca 1140 atatgttacc tcatttgtag ctggcacata ttctttcact ttcaatagat ttctaaattt 1200 agtcaccaac ccaaatcccg atttcaggat gtttgtcgag tcatgcaaac gtttgcgcat 1260 aatgaaagga tccgaagcaa ttggacttgg taagttttct tttctgttcg tttctataag 1320 tggctctttt ctgtttttcc aataatgctc gtgttttttt ttcagctcca agagcaatgg 1380 agaagttcaa gaacagatca tga 1403 5 21 DNA Artificial Sequence synthetic oligonucleotide 5 catattctga tttaagacat a 21 6 21 DNA Artificial Sequence synthetic oligonucleotide 6 aatcaatgca tattgtcctc t 21 7 21 DNA Artificial Sequence synthetic oligonucleotide 7 ttatggctaa tcagaagagc g 21 8 21 DNA Artificial Sequence synthetic oligonucleotide 8 tattctctaa acaaaaaaaa c 21 9 21 DNA Artificial Sequence synthetic oligonucleotide 9 aattcgactt ctgggtactc a 21 10 21 DNA Artificial Sequence synthetic oligonucleotide 10 aaattaagtc cctcaagctg g 21 11 21 DNA Artificial Sequence synthetic oligonucleotide 11 actctgaatt tgtagaaaga a 21 12 21 DNA Artificial Sequence synthetic oligonucleotide 12 gaagatgatt ttgttgccat a 21 13 21 DNA Artificial Sequence synthetic oligonucleotide 13 aagatgggga gctggaatat c 21 14 21 DNA Artificial Sequence synthetic oligonucleotide 14 ggctcaacac cctctagcat a 21 15 21 DNA Artificial Sequence synthetic oligonucleotide 15 catccatacc agcttgaggg a 21 16 21 DNA Artificial Sequence synthetic oligonucleotide 16 caagtttgat gtcctcctca c 21 17 21 DNA Artificial Sequence synthetic oligonucleotide 17 acatgccccc caaaaaggag c 21 18 21 DNA Artificial Sequence synthetic oligonucleotide 18 ccatcaattc gctcgtactg c 21 19 21 DNA Artificial Sequence synthetic oligonucleotide 19 atgtgctgaa actgtgtgta c 21 20 21 DNA Artificial Sequence synthetic oligonucleotide 20 ccattgcttt tgctgacgca t 21 21 21 DNA Artificial Sequence synthetic oligonucleotide 21 ttcgatagcc aaccacagtc t 21 22 21 DNA Artificial Sequence synthetic oligonucleotide 22 ggcatgcaat atgggtggcg t 21 23 21 DNA Artificial Sequence synthetic oligonucleotide 23 tcaggtatgg atcaaaggag c 21 24 21 DNA Artificial Sequence synthetic oligonucleotide 24 ctcccctcac cttccatcaa c 21 25 21 DNA Artificial Sequence synthetic oligonucleotide 25 gtgcacaatc ttgtcaaatc a 21 26 21 DNA Artificial Sequence synthetic oligonucleotide 26 gaggcacaga gagtcgctgc t 21 27 21 DNA Artificial Sequence synthetic oligonucleotide 27 tatacattgg tttggtctgc c 21 28 21 DNA Artificial Sequence synthetic oligonucleotide 28 gtagggatag atgatgagcc a 21 29 21 DNA Artificial Sequence synthetic oligonucleotide 29 ccccgatgca tctaaattat c 21 30 21 DNA Artificial Sequence synthetic oligonucleotide 30 actagttcag gagaaggtga g 21 31 21 DNA Artificial Sequence synthetic oligonucleotide 31 acatgcagag acgacttgtt g 21 32 21 DNA Artificial Sequence synthetic oligonucleotide 32 cggacttcat cgaacctatt c 21 

What is claimed is:
 1. An Arabidopsis thaliana double mutant ssl2 slr having a mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, obtained by: treating an Arabidopsis thaliana slr dominant mutant (FERM BP-8385), which has no lateral roots, with a mutagen; preparing plants of the next generation of the mutagen-treated slr dominant mutant; and selecting a plant that basically preserves phenotypes of the slr dominant mutant but has lateral roots from the plants of the next generation.
 2. An Arabidopsis thaliana double mutant ssl2 slr, which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant (FERM BP-8385) that has no lateral roots, due to an additional mutation of at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant.
 3. An Arabidopsis thaliana double mutant ssl2 slr, which has recovered the capability of lateral root formation in an Arabidopsis thaliana slr dominant mutant (FERM BP-8385) that has no lateral roots, due to an additional mutation of the SSL2 genomic gene shown in SEQ ID NO: 3 in the slr dominant mutant, wherein the additional mutation is selected from the group consisting of the following (A) to (D): (A) a mutation in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; (B) a mutation in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; (C) a mutation in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; and (D) a mutation in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”.
 4. A mutant gene having a mutation in at least one base of the SSL2 gene (cDNA) shown in SEQ ID NO: 1, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.
 5. A mutant gene having a mutation in at least one base of the SSL2 genomic gene shown in SEQ ID NO: 3, whose expression enables a phenotype of a mutant that has no lateral roots to be recovered.
 6. A mutant gene of the SSL2 gene (cDNA) selected from the group consisting of the following (a) to (c): (a) a mutant gene in which the 566th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”; (b) a mutant gene in which the 1005th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”; and (c) a mutant gene in which the 901th base “G” of the SSL2 gene (cDNA) shown in SEQ ID NO: 1 has been substituted with “A”.
 7. A mutant gene selected from the group consisting of the following (d) to (g): (d) a mutant gene in which the 852th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; (e) a mutant gene in which the 4734th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; (f) a mutant gene in which the 1757th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”; and (g) a mutant gene in which the 1546th base “G” of the SSL2 genomic gene shown in SEQ ID NO: 3 has been substituted with “A”.
 8. A protein selected from the group consisting of the following (a) and (b): (a) a protein comprising the amino acid sequence of SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked; and (b) a protein comprising an amino acid sequence of SEQ ID NO: 2, in which one or a few amino acids of the amino acid sequence have been deleted, substituted and/or added and which has a function of maintaining a mutation whereby lateral root formation is blocked.
 9. A gene encoding a protein selected from the group consisting of the following (a) and (b): (a) a protein comprising the amino acid sequence of SEQ ID NO: 2 and having a function of maintaining a mutation whereby lateral root formation is blocked; and (b) a protein comprising an amino acid sequence of SEQ ID NO: 2, in which one or a few amino acids of the amino acid sequence have been deleted, substituted and/or added and which has a function of maintaining a mutation whereby lateral root formation is blocked.
 10. A gene selected from the group consisting of the following (c) or (d): (c) a gene comprising the DNA sequence of SEQ ID NO: 1 and encoding a protein having a function of maintaining a mutation whereby lateral root formation is blocked; and (d) a gene comprising a DNA sequence of SEQ ID NO: 1, in which one or a few bases of the DNA sequence have been deleted, substituted and/or added and which encodes a protein having a function of maintaining a mutation whereby lateral root formation is blocked. 